Enhancement of late successional plants on ex-arable land by soil inoculations

Restoration of species-rich grasslands on ex-arable land can help the conservation of biodiversity but faces three big challenges: absence of target plant propagules, high residual soil fertility and restoration of soil communities. Seed additions and top soil removal can solve some of these constraints, but restoring beneficial biotic soil conditions remains a challenge. Here we test the hypotheses that inoculation of soil from late secondary succession grasslands in arable receptor soil enhances performance of late successional plants, especially after top soil removal but pending on the added dose. To test this we grew mixtures of late successional plants in arable top (organic) soil or in underlying mineral soil mixed with donor soil in small or large proportions. Donor soils were collected from different grasslands that had been under restoration for 5 to 41 years, or from semi-natural grassland that has not been used intensively. Donor soil addition, especially when collected from older restoration sites, increased plant community biomass without altering its evenness. In contrast, addition of soil from semi-natural grassland promoted plant community evenness, and hence its diversity, but reduced community biomass. Effects of donor soil additions were stronger in mineral than in organic soil and larger with bigger proportions added. The variation in plant community composition was explained best by the abundances of nematodes, ergosterol concentration and soil pH. We show that in controlled conditions inoculation of soil from secondary succession grassland into ex-arable land can strongly promote target plant species, and that the role of soil biota in promoting target plant species is greatest when added after top soil removal. Together our results point out that transplantation of later secondary succession soil can promote grassland restoration on ex-arable land.     

The effects of three years of fencing enclosure on soil seed banks and the relationship with above-ground vegetation of degraded alpine grasslands of the Tibetan plateau

Background and Aims

Fenced enclosures have become an important method for re-establishing degraded grassland on the Tibetan plateau, and examination of soil seed banks may provide useful insights to understanding the effects and mechanisms of fencing enclosure on the restoration.

Methods

An investigation was conducted into the effects of enclosure for 3 years on the soil seed banks of degraded natural and sown grasslands at eight study sites. Species composition, soil seed bank density and the relationships with above-ground vegetation were analysed based on 4800 soil core samples and counting of seeds extracted from soil samples.

Results

After 3 years of fencing enclosure, soil seed banks differed between the different communities across the study sites. Species numbers and seed density in soil seed banks decreased from natural grassland to sown grassland, with most seeds occurring in the upper 5 cm soil layer. In these alpine grasslands, relatively few species produced high numbers of seeds, although their occurrence across the eight study sites was variable. Total vegetation cover increased with enclosure due to the colonization capacity of the vegetation rather than soil seed banks.

Conclusions

This study provided evidence that soil seed banks do not play an important role in the restoration of degraded alpine grassland when using fencing enclosures. Further studies conducted over longer periods are needed to address this subject.     

Species introduction in restoration projects – Evaluation of different techniques for the establishment of semi-natural grasslands in Central and Northwestern Europe

During recent decades, many studies have shown that the successful restoration of species-rich grasslands is often seed-limited because of depleted seed banks and limited seed dispersal in modern fragmented landscapes. In Europe, commercial seed mixtures, which are widely used for restoration measures, mostly consist of species and varieties of non-local provenance. The regional biodiversity of a given landscape, however, can be preserved only when seeds or plants of local provenance are used in restoration projects. Furthermore, the transfer of suitable target species of local provenance can strongly enhance restoration success.We review and evaluate the success of currently used near-natural methods for the introduction of target plant species (e.g. seeding of site-specific seed mixtures, transfer of fresh seed-containing hay, vacuum harvesting, transfer of turves or seed-containing soil) on restoration sites, ranging from dry and mesic meadows to floodplain grasslands and fens. Own data combined with literature findings show species establishment rates during the initial phase as well as the persistence of target species during long-term vegetation development on restoration sites.In conclusion, our review indicates that seed limitation can be overcome successfully by most of the reviewed measures for species introduction. The establishment of species-rich grasslands is most successful when seeds, seed-containing plant material or soil are spread on bare soil of ex-arable fields after tilling or topsoil removal, or on raw soils, e.g. in mined areas. In species-poor grasslands without soil disturbance and on older ex-arable fields with dense weed vegetation, final transfer rates were the lowest. For future restoration projects, suitable measures have to be chosen carefully from case to case as they differ considerably in costs and logistic effort. Long-term prospects for restored grassland are especially good when management can be incorporated in agricultural systems.     

The seed bank in an estonian calcareous grassland: Comparison of different successional stages

Seed bank species-composition and seed-density were determined in a successional calcareous (alvar) grassland in western Estonia. Three similar study areas were chosen to compare two different successional stages: open alvar grassland and overgrown areas with young pine forest 30 to 40 years old. In both successional stages, the centre and the edge of a relatively uniform stand were examined. Fifteen soil samples (7 cm in diameter, 5 cm deep) were taken from each of twelve sampling sites. The seedling emergence method was used to estimate seeds in the soil samples. A total of 69 species were detected in the seed bank, of which 18 did not occur in the vegetation. Eighty-nine taxa were recorded in the vegetation and of these 38 were not detected in the seed bank. Fifty-one species occurred both in the seed bank and in the vegetation. The three most abundant taxa in the seed bank wereCarex tomentosa, Linum catharticum andPlantago media, which together made up 49% of the seedlings recorded. Differences in the species compositions of seed bank samples from grassland and forest sites were negligible, although the species richness per area of the above-ground vegetation was significantly higher in the open grassland. The only species tending to be lost from forest site vegetation but still occurring in the forest soil seed bank wereArenaria serpylifolia, Cerastium fontanum andLinum catharticum. About half of all the emerged species from all samples belonged to the transient or short-term persistent seed bank. In the grassland sites there were more species which belonged to the transient seed bank than in the forest sites, where the seed bank contained more short-term persistent type seeds. The seed density was significantly higher in forest sites and lower in grassland sites, which may be explained by the better germination conditions in well-illuminated communities. On the basis of the current study it might be assumed that the soil seed banks of overgrown alvar grasslands which include young pine forests can play a certain role in grassland restoration management.     

Multivariate analysis of the response of overgrown semi-natural calcareous grasslands to restorative shrub cutting

Monitoring and evaluation of restoration management often suffers from poor sampling design and a lack of statistical rigour, seldom considering the spatial and temporal variability of habitat. We tested the effectiveness of shrub cutting on the restoration of calcareous grasslands, using a 4-year Before–After/Reference-Control-Impact (BARCI) project design with replicated patches of habitat. Departure from the control sites and convergence with the reference sites were analysed using the Principal Response Curves (PRC) method.The structure of communities was compared 1 year before and 3 years after management on control scrubland, restored ex-arable land, restored and reference grassland. Results show that shrub cutting did not suffice to restore the community composition of the semi-natural calcareous grasslands. The restored ex-arable areas maintained a community structure extremely similar to the control scrubland.The restored grassland, on the other hand, deviated from the control and slightly converged to the reference grassland, though not significantly. The dominant scrub species that was cut, Prunus spinosa, showed higher cover values compared to the reference grassland in any of the treatments, even the first year after cutting. Species typical of xeric and/or calcareous grassland were more abundant in the reference than in the restored grassland, while arable and ruderal species were more frequent in the restored sites.This study has demonstrated that the BARCI approach is a powerful tool for the evaluation of restoration management, as it was possible to evaluate not only departure from the unmanaged control, but also convergence with the reference community. The PRC method provided a comprehensive overview of the response of the 153 species involved in the study.Our results also indicate that the second PRC should be considered, when significant, to not exclude important information from the assessment.     

Disturbance regimes as determinants of seed banks in coastal dune vegetation of the southeastern Cape

Soil-stored seed banks of grassland, fynbos and thicket, all growing on calcareous dunes and each subject to different disturbance regimes, were examined. Seed banks were determined from counts of germinants from 50 soil cores from each type. Aboveground estimates of plant species cover in 10 1-m² plots were used in determining vegetation/seed bank similarities. There was no evidence for seed bank densities to be markedly higher in the most frequently disturbed community (grassland -4273 seeds/m²) than the least disturbed community (thicket - 3417 seeds/m²). Highest similarity between seed bank and above-ground vegetation composition in terms of species and growth form/life-span classes was recorded for grassland (CC = 50%). Lowest similarity (CC = 13%) was found in the less frequently disturbed thicket where no seeds of climax trees were recorded in the seed bank. A fynbos community on a north-facing (warm, dry) slope had intermediate-sized seed banks (1683 seeds/m²) with intermediate vegetation/seed bank similarity (CC = 46%). However, on the south-facing slope, which has a large post-fire ephemeral herb component, seed banks were larger (4518 seeds/m²) but less similar to above-ground vegetation (CC = 39%o). Ordination (DCA) of vegetation data from the four communities was different from an ordination of their seed bank data. Fynbos shrub species were absent from seed banks of both grassland and thicket, even though secondary succession proceeds from grassland, through fynbos to thicket. Their seed banks appear less persistent than those of European heath or Californian chaparral shrubs.     

Soil seed-bank composition reveals the land-use history of calcareous grasslands

We compared soil seed banks and vegetation of recent (established on abandoned arable fields) and ancient (continuously managed as pastures at least since 1830) calcareous grasslands if there is any impact of former arable field use. The study was carried out in two regions of Southern Germany with well-preserved dry grassland vegetation: the western Jurassic mountains (Kaltes Feld) and the climatically drier eastern part of Southern Germany (Kallmünz).Total number of species in the seed bank was similar in both regions, but species composition partly differed, reflecting phytogeographical differences between the regions. The total number of emerged seedlings showed a large disparity (5457 compared to 2523 seedlings/m² in Kaltes Feld and Kallmünz, respectively).Though there were differences in seed bank composition and size, we found a uniform pattern of plant traits (affiliation to phytosociological groups, Raunkiaer plant life-forms and seed longevity), which depended on the age of the grassland.The main conclusion is that seed banks in contemporary calcareous grasslands still reflect the history of former land use – in this case arable cultivation, even though it occurred a long time ago (up to 150 years). Indicators of former arable fields are germinable seeds of weeds which have persisted in the soil to the present. By contrast, weedy species are completely absent from the seed banks of ancient grasslands. Soil seed banks of recent grasslands may be of substantial conservation importance because they may store seeds of rare and endangered weed species such as Kickxia spuria, Silene noctiflora and Stachys annua, the majority of which have already gone extinct from the current vegetation of the study sites.     

Intraspecific trait variation and allocation strategies of calcareous grassland species: Results from a restoration experiment

Intra- and interspecific trait variation express the response of plants dealing with different environmental conditions. We measured root and leaf traits on 14 species of calcareous grasslands in a restoration experiment. We aimed at identifying intraspecific differences in biomass allocation and functional plant traits under contrasting soil conditions by comparing plants growing in ancient grassland and two restored grasslands on ex-arable land, one of them with topsoil removal. Relative importance of trait variation within and among species, and among site was assessed by variance partitioning. Interspecific variation was more important than intraspecific variation, but the contribution of the latter to total variation was considerable, especially for specific leaf area. Changes in soil properties due to topsoil removal resulted in lower values of plant height, specific leaf area and specific root length compared to the control (ancient grassland). Soil fertility found in the treatment without top soil removal did not affect plant plasticity compared to the control. The study species showed two allocation strategies in relation to resource stress, while the responses of individual traits to the soil treatments were consistent across species. We conclude that caution must be taken when using mean trait values for plastic species or when working with environmental gradients.     

Interactions between plant and insect diversity in the restoration of lowland calcareous grasslands in southern Britain

Abstract. The lowland calcareous grasslands of northwestern Europe are highly prized by ecologists and conservationists as a result of the diversity of their plant and invertebrate communities. Large areas of such grasslands have been lost this century as a result of changes in agricultural land use. Recent changes in agricultural policies, in particular the introduction of agri-environmental incentive schemes, have resulted in an increasing area being managed for the restoration of these communities. This paper reviews the management techniques employed in the restoration of lowland calcareous grasslands and the factors that govern their success. Constraints on the enhancement of the plant diversity of restoration sites include high soil fertility and the presence of undesirable species in the soil seed bank. However, it is thought that the primary constraint is the availability of propagules from which new populations can be established. Similarly, the dispersal mode and ability of insect species is likely to be the major factor limiting the enhancement of insect diversity. Evaluation of the success of restoration management usually involves monitoring changes in the plant community. However, as a result of their short life-cycles and sensitivity to small-scale environmental conditions, insects may respond more rapidly to changes resulting from restoration management and therefore provide better indicators of success. With the exception of a few high-profile butterfly species, the use of insects as indicator taxa has largely been neglected in terrestrial systems. This paper illustrates their potential use with reference to lowland calcareous grasslands in southern Britain.     

荒漠草原土壤种子库对草地管理方式的响应

以宁夏中部不同封育年限(封育1a、3a、5a和7a)、不同放牧方式(中等强度自由放牧、中等强度4区轮牧)、补播改良(2006年补播沙打旺Astragalus adsurgens和紫花苜蓿Alfalfa stiva)及未封育等管理方式下的荒漠草原为研究对象,采用空间梯度代替时间梯度法,于2012年3月下旬用样线法采集土壤样品,采用温室萌发法,统计土壤种子库物种组成和种子数,研究了不同草地管理方式下的种子库与地上植被相似性、多样性等特征。结果表明:1)宁夏荒漠草原土壤种子库中共出现14种植物,分属6科14属,除牛枝子(Lespedeza potanimii)为半灌木外其余均为草本植物。多年生植物占土壤种子库植物总数的65.3%,一年生植物占土壤种子库植物总数的35.7%;多年生植物种子数量随封育年限增加而表现为下降趋势,封育1a和未封育草地最多;2)不同管理方式下土壤种子库和地上植被的共有物种数为2—6种,封育时间延长导致土壤种子库物种与地上植被相似性下降,土壤种子库和地上植被共有物种数有减少趋势,自由放牧降低了土壤种子库和地上植被的共有物种数,补播恢复的草地土壤种子库中未出现补播物种;3)种子库物种多样性与管理方式无明显的相关性;种子库物种多样性指数4区轮牧草地最高,封育7a草地最低,物种丰富度指数封育1a草地最高,均匀度指数自由放牧草地最高,补播草地最低。若荒漠草原仅依靠土壤种子库恢复自然植被,则不建议封育时间过久,需间以适当的干扰(放牧);补播外来种未对荒漠草原土壤种子库产生贡献,反而对乡土种产生了一定限制。     

Rapid restoration of a species-rich ecosystem assessed from soil and vegetation indicators: The case of calcareous grasslands restored from forest stands

Calcareous grasslands have long been recognized as biodiversity hotspots in Europe. However, in recent decades these ecosystems have seen rapid decline. In Belgium, more than 100 ha of calcareous grasslands have been restored from oak coppices and pine forests since the 1990s. The aim of the present study was to provide a quantitative assessment of the success of these restoration efforts, using two sets of indicators: one related to soil conditions, the other related to vascular plant communities. Soil conditions were evaluated by comparing soil samples from pre-restoration forest stands, restored grasslands (3-age classes: 2–4 years; 5–8 years, and 10–15 years) and reference grasslands. The analysis revealed no significant differences in soil N, P, and K contents between pre-restoration forests and restored and reference grasslands. We observed a decrease in the mineralization rate indicators in both pre-restoration forests and recent grassland restorations, which was resorbed in older restorations. Floristic surveys revealed that plant species composition of older restorations was most like reference grasslands. However, some differences in species composition persisted after 15 years. Moreover, a few rare species did not colonize restored grasslands despite a close seed source. Non-recolonization by a set of species expected on calcareous grasslands may be due to dispersal limitation and higher cover by native invasive grasses in restored parcels. These results were discussed in term of implications for management.     

Plant community composition and attributes reveal conservation implications for newly created grassland on capped landfill sites

Biodiversity conservation through restoring degraded habitats or creating new habitats is advocated in the UK Biodiversity Action Plan. Restored grasslands on capped landfill sites are semi-natural habitats that could potentially support a wide range of plant communities. However, it is unknown whether these re-created habitats represent a significant resource in terms of biodiversity conservation. The UK National Vegetation Classification (NVC) was used to study these communities on nine restored capped landfill sites together with paired reference sites in the East Midlands region of the UK during 2007. Plant species data were collected by random quadrats along two 100 m transects from each site. The effects of restoration were investigated by examining plant attributes and Ellenberg indicators on restored landfill sites in comparison to paired reference sites. A total of 170 plant species were found from both sets of sites. There were no significant differences for most of the plant attributes between restored landfill sites and reference sites, though reference sites had significantly higher mean frequencies of native plants, nationally decreasing species and perennial species. In total 26 broad NVC community types were identified of which more than 70% fell within mesotrophic grassland (MG). The diversity of NVC communities confirms that underlying environmental factors such as soil fertility are important, dictating the type of plant communities that exist. It is recommended that management of these capped landfill sites should be targeted towards specific NVC community types to meet conservation targets.     

Heat shock increases the reliability of a temperate calcareous grassland seed bank study

Reliable assessment of the density and species richness of the viable seeds in the soil is essential to estimate the probability of successful restoration of a particular plant community. Since temperate calcareous grasslands contain several thermophilous species typical of fire prone habitats, heat shock can be expected to break dormancy of target species for calcareous grassland restoration. We tested the effect of heat shock on the estimated density and species richness of seeds in soil samples. Heat shock treatments between 70 and 110°C resulted in both a higher estimated seed density and estimated species richness, in particular of specialist calcareous grassland species. Moreover, 25 species germinated exclusively from the heat-treated samples. This indicates that several species would have been missed by handling soil seed bank samples following the standard germination protocol, resulting in an underestimation of the restoration potential.     

The roles of seed mass and persistent seed banks in gap colonisation in grassland

Gaps in grassland created by animals are often sites for species regeneration. The persistent seed banks of ant-hills and surrounding soils in a calcareous grassland in southern Britain were compared and analysed in terms of seed mass and longevity. The relative abundance of species on ant-hills compared to the pasture was highly correlated with seed abundance in ant-hill soil. The abundance of plant species on ant-hills compared to the pasture was significantly negatively correlated with seed mass, and positively correlated with seed longevity. These results suggest that germination from the seed bank in ant-hill soils, as well as limited seed dispersal and vegetative invasion, contributes to the distinctive ant-hill vegetation. The same may apply to other gaps created by animals in grasslands.     

科尔沁沙质草地放牧和围封条件下的土壤种子库

该文研究了科尔沁沙质草地在放牧和围封条件下土壤种子库密度、组成及其与地上植被的关系。结果表明:1)放牧草地植物种数22种,围封草地植物种数30种,围封使土壤种子库植物种数增加了36%;2)放牧草地土壤种子库密度为16 149±1 900有效种子数·m^-2, 围封草地土壤种子库密度为20 657±3 342有效种子数·m^-2,比放牧草地增加了28%。放牧和围封草地种子库组成密度均以一年生植物为主(分别占99%和98%的比例),多年生植物所占的比例很小;3)放牧草地种子库的Shannon-Wiener指数和丰富度指数分别为0.836 3和4.954 9,明显小于围封草地的0.968 2和7.226 0,表明自由放牧导致物种多样性下降;4)放牧和围封草地土壤种子库密度与地上植被密度均存在显著的相关性(p<0.001)。表明了随着土壤种子库密度的增加, 地上植被密度随之增加,放牧草地地上植被密度78%的变异可归结为土壤种子库密度的变异, 而围封草地地上植被密度58%的变异可由土壤种子库密度的变异来解释。     

Are Ants Useful Indicators of Restoration Success in Temperate Grasslands?

Assessments of restoration are usually made through vegetation community surveys, leaving much of the ecosystem underexamined. Invertebrates, and ants in particular, are good candidates for restoration evaluation because they are sensitive to environmental change and are particularly important in ecosystem functioning. The considerable resources currently employed in restoring calcareous grassland on ex‐arable land mean that it is important to gather as much information as possible on how ecosystems change through restoration. We compared ant communities from 40 ex‐arable sites where some form of restoration work had been implemented between 2 and 60 years previously, with 40 paired reference sites of good quality calcareous grassland with no history of improvement or cultivation. A total of 11 ant species were found, but only two of these were found to be significantly different in abundance between restoration and reference sites: Myrmica sabuleti was more likely to be present in reference sites, whereas Lasius niger was more likely to be found in restoration sites. Myrmica sabuleti abundance was significantly positively correlated with age of restoration sites. The potential number of ant species found in temperate grasslands is small, limiting the information their assemblages can provide about ecosystem change. However, M. sabuleti is a good indicator species for calcareous grassland restoration success and, alongside information from the plant community, could increase the confidence with which restoration success is judged. We found the survey to be quick and simple to carry out and recommend its use.     

兰坪铅锌矿区植被恢复初期土壤种子库与地上植被关系的研究

尾矿废弃地是一种极端的生态系统,其植被恢复的研究将丰富传统的生态学理论。该研究通过野外植被调查与室内萌发实验相结合的方法,探讨了兰坪铅锌矿区植被恢复初期不同群落类型地上植被、土壤种子库及其相互关系。结果显示:（1）与对照群落（云南松林、高山栎灌丛）相比,尾矿区恢复期各群落（早熟禾人工草地、魁蒿群落、马桑灌丛）地上植被及土壤种子库的物种数、物种多样性均较低。（2）植被恢复时间较短的2个群落（魁蒿群落、人工草地）土壤种子库较地上植被物种多样性高。（3）尾矿区恢复期各群落地上植被及土壤种子库的优势种均主要由风播、种子繁殖的植物组成,菊科、禾本科占较大比例,这些植物在尾矿区植被恢复初期起重要作用。（4）尾矿区恢复期各群落土壤种子库与地上植被的物种相似性较高,各群落之间地上植被及土壤种子库的相似性则较低。研究表明,尾矿区恢复初期土壤种子库与地上植被紧密联系,群落改造方式、恢复时间对土壤种子库具有重要影响。     

Sowing different mixtures in dry acidic grassland produced priority effects of varying strength

Several longer-term assembly studies on ex-arable land have found that species that arrive first at a disturbed site can play a key role in the further development of the community and that this priority effect influences aboveground productivity, species diversity and stability of the grassland communities that develop. Restoration of nutrient poor, species rich grasslands is often limited by seed dispersal as well as the accessibility of suitable microsites for establishment. Sowing species (i.e. creating priority effects for further assembly) may help overcome such dispersal barriers, but the potential of using priority effects for restoration has not been tested in this type of dry grassland. We tested the hypothesis that sowing two different seed mixtures used for dry acidic grassland restoration onto a sandy substrate (which formed an equivalent to a primary succession) would create priority effects, and that these priority effects would be sustained over a number of years. We followed community assembly and measured aboveground productivity for four years after sowing. We found that priority effects caused by sowing of differently diverse mixtures did also occur in dry acidic grassland habitat, but that how persistent they were over time depended on the response variable considered. Priority effects on species number were not as strong as found in previous ex-arable land studies, whereas priority effects for aboveground productivity were still visible after 4 years. In addition, functional composition of the community still reflected the composition of the seed mixtures 4 years later. Our results suggest that priority effects can occur in nutrient-poor dry acidic grassland but in contrast to more nutrient-rich sites the breadth of responses affected may not be as wide.     

Vegetation and soil seed bank in a 23-year grazing exclusion chronosequence in a Mediterranean dry grassland

Long-term grazing shaped plant diversity in dry Mediterranean grasslands. Abandonment of grazing affects plant diversity especially in the northern Mediterranean. Considerable efforts are, therefore, under way for grassland conservation and restoration. Yet, we do not know at which temporal scales impacts of grazing abandonment appear and in particular how soil seed banks evolve after longer grazing abandonment. Here, we provide detailed data from one of the very few long-term experiments available. These experiments provide data for up to 23 years (1982–2005) of grazing exclusion built in 1982, 1989, 2000 and 2001. Grazing exclusion decreased species richness, modified vegetation structure and changed soil parameters. Decline in species richness appears in communities that experienced 16 and 23 years of grazing exclusion. Only four to nine plant species of this Mediterranean grassland built persistent soil seed banks appearing after grazing exclusion, compared to 40–50 species in the established vegetation of grazed plots. Hence, similarity between vegetation and soil seed bank decreased with time of grazing exclusion. Even 23 years after abandonment, no woody plants colonised the experiments. We conclude that vegetation will recover fast from grazing abandonment in the short-term. Nevertheless, longer abandonment will impact diversity due to reduced soil seed banks.     

Soil Organism and Plant Introductions in Restoration of Species-Rich Grassland Communities

Soil organisms can strongly affect competitive interactions and successional replacements of grassland plant species. However, introduction of whole soil communities as management strategy in grassland restoration has received little experimental testing. In a 5-year field experiment at a topsoil-removed ex-arable site ( receptor site ), we tested effects of (1) spreading hay and soil, independently or combined, and (2) transplanting intact turfs on plant and soil nematode community development. Material for the treatments was obtained from later successional, species-rich grassland ( donor site ). Spreading hay affected plant community composition, whereas spreading soil did not have additional effects. Plant species composition of transplanted turfs became less similar to that in the donor site. Moreover, most plants did not expand into the receiving plots. Soil spreading and turf transplantation did not affect soil nematode community composition. Unfavorable soil conditions (e.g., low organic matter content and seasonal fluctuations in water level) at the receptor site may have limited plant and nematode survival in the turfs and may have precluded successful establishment outside the turfs. We conclude that introduction of later successional soil organisms into a topsoil-removed soil did not facilitate the establishment of later successional plants, probably because of the "mismatch" in abiotic soil conditions between the donor and the receptor site. Further research should focus on the required conditions for establishment of soil organisms at restoration sites in order to make use of their contribution to grassland restoration. We propose that introduction of organisms from "intermediate" stages will be more effective as management strategy than introduction of organisms from "target" stages.